Differential and reversible responses of common fen meadow species to abandonment

Abstract. We studied the effects of abandonment on two common fen plant species. In mown and a chronosequence of abandoned fen meadows spanning 35 yr, we measured fitness traits of the sedge Carex davalliana and the forb Succisa pratensis. Cessation of mowing had little effect on fitness traits and seed production of C. davalliana, but seedling density decreased more than threefold. Population density of S. pratensis decreased with increasing community biomass, but was not affected by the cessation of mowing. However, flowering frequency increased threefold and seed production was 20% higher in fallow meadows. Consequently, seedling density of S. pratensis increased nearly threefold after abandonment. However, these changes were not dependent on the age of the fallow. In a common garden and germination experiment, we found no differences in either species between plants from fallows and mown fen meadows, except for the height of the flowering stalk of S. pratensis. The combined results from the common garden experiment and the field studies indicate that changes in fitness traits observed in fallows were mostly phenotypic and likely to be reversible. If other species react in similar ways, there is a high potential for re‐establishing traditional fen meadow communities from fallows by mowing.     

Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

Seed deposition in drift lines: Opportunity or hazard for species establishment?

Arrival at suitable sites for germination and establishment is crucial for hydrochorous plants. Seed deposition in riparian landscapes occurs mainly within litter in drift lines. We investigated whether conditions within drift lines are suitable for seed survival and germination and whether drift lines act as seed sources or sinks for species diversity in riparian fen grasslands.We analysed drift litter in a small river valley in Northern Germany (Upper Eider valley) after typical late winter floods in 2000, 2002 and 2004. The content of viable seeds in drift lines was studied via a germination trial with drift litter samples taken from five fen grassland sites representative of the vegetation of floodplain. Further, seedling recruitment of both abandoned and moderately grazed fen grasslands affected by drift line deposition was compared with equivalent sites without drift lines. Additionally, the decline of viable seeds after the deposition of drift litter was studied. Finally, the relative importance of seed-containing drift litter and microsite availability for seedling recruitment was investigated in a field experiment with the factors disturbance (mowing, large gaps, small gaps, undisturbed control) and addition of (seed-containing) drift litter (yes, no) in an abandoned riparian fen.Seedlings which recruited from drift litter collected at different sites differed in species composition. A nearly 50% reduction of the number of viable seeds in drift litter was observed already during 2 weeks after deposition. Naturally deposited drift lines had a positive effect on the number of seedlings, although an increase of seedling recruitment in areas with drift litter was only observed at the grazed site. Adding drift litter reduced the number of seedlings in the field experiment, while creating gaps enhanced seedling recruitment.These results indicate that in riparian fen grasslands, drift lines can affect species diversity either negatively by hampering germination and/or establishment or positively by increasing seed availability. As the studied river system is comparatively species poor and flooding extension is limited, the role of drift lines as seed source might be of minor importance in the study area.     

Does Germination Success Differ with Respect to Seed Mass and Germination Season? Experimental Testing of Plant Functional Trait Responses to Grassland Management

BACKGROUND AND AIMS: Plant functional trait responses to processes such as grassland management have been analysed frequently; however, the scaling-up from individual traits to the outcomes of vegetation dynamics has seldom been tested. In this experiment, germination success was studied with respect to the relationships between grassland management (mowing and grazing), as well as abandonment, and two traits that are relevant for seedling recruitment: seed mass and germination season. On the basis of discussions in the literature and indirect trait analyses in our previous studies, the following hypotheses are proposed: (1) with respect to seed mass, mowing and grazing favour the germination of small seeds, whereas after abandonment the germination success of larger seeds is higher; and (2) with respect to germination season, mowing and grazing favour autumn-germinating seeds, whereas succession promotes spring-germinating seeds. METHODS: The germination experiment took place in a semi-natural, dry grassland in north-east Germany. Seeds of eight herbaceous species that differ with respect to seed mass and germination season were sown in mown, grazed and abandoned plots. Germination success was documented during the following year. KEY RESULTS AND CONCLUSIONS: Contrary to the hypothesis, germination of small seeds was not promoted by mowing or grazing and they germinated relatively more often than expected in the abandoned plots. A relationship between abandonment and gaps of bare soil below the vegetation cover that favour germination of small seeds was likely, but could not be proved statistically. It is possible that the small seeds suffered less from predation. Mowing favoured autumn germination, which could be explained by the removal of biomass in late summer. Contrary to our expectation, there was relatively more spring germination after grazing than after mowing, yet vegetation height was smallest in spring. Generally, germination season was found to be related to the temporal occurrence of favourable light conditions.     

Restorative mowing on an abandoned semi‐natural meadow: short‐term and predicted long‐term effects

Abstract. When management, in the form of cattle grazing and mowing, ceases the abundance of competitively superior plant species tends to increase in abandoned semi‐natural meadows. Litter accumulation elevates the soil nutrient levels and hinders seedling recruitment. We surveyed changes in plant cover and species composition of a formerly grazed meadow in permanent plots for six years. Some plots were unmown, while others were mown and raked annually in August. The cover of grasses decreased and herb cover remained unchanged regardless of the treatment. Mowing and raking significantly reduced litter accumulation and increased the number of ground layer species. The expected long‐term effects of abandonment and restorative mowing were studied by calculating the transition probabilities for unmown and mown plots and simulating the course of succession as projected by the transition matrices. During a simulation period of 30 yr, abandonment led to (1) a decrease in the cover of small herbs, (2) a slight increase in the cover of tall herbs and (3) a slight decrease in the cover of grasses. In contrast, the cover of small herbs on the mown plots remained unchanged or slightly increased during the course of simulation. These results suggest that mowing late in the season is primarily a management tool for the maintenance of the existing species diversity and composition. However, it may not be an effective restorative tool to induce overall changes in the resident vegetation of abandoned grass‐dominated meadows. Grazing or mowing early in the season may be more effective in this respect. Consequently, mowing early or, alternatively, late in the season may provide management strategies for the maintenance and restoration of species diversity, respectively.     

Factors Affecting Seed Germination and Seedling Establishment of Fen-Meadow Species

Availability of seeds and provision of “safe sites” for seedling recruitment are essential for successful restoration of seminatural grassland communities. Inability to provide species‐specific conditions for seedling recruitment appears to be a major factor limiting establishment of fen‐meadow species on restoration sites. This contention was tested in the field for both germination and establishment conditions for a selection of fen‐meadow species. A Cirsio‐Molinietum fen meadow and an agriculturally semi‐improved species‐poor grass dominated rush pasture were used. Seeds of Carex ovalis, Cirsium dissectum, Molinia caerulea, Succisa pratensis, and Holcus lanatus were sown onto treatments comprising either irrigation or no irrigation, presence or absence of existing vegetation canopy, and presence or absence of soil disturbance. Germination of all except H. lanatus was higher in the fen meadow than in the rush pasture. The fen‐meadow site was less susceptible to drought, provided more light to the seed environment, and showed a stronger day–night variation in relative humidity compared with the rush pasture. All the fen‐meadow species responded strongly to the experimental treatments, whereas H. lanatus showed only a small response. Soil disturbance was the major factor that increased germination. Removal of the vegetation canopy improved germination only in S. pratensis. Conditions affecting survival of seedlings were different from those affecting seed germination. Seedling survival was greater on the fen‐meadow site than on the rush pasture. Canopy presence was the major factor that reduced seedling survival. Few seedlings survived in the presence of the rush pasture canopy. Irrigation and soil disturbance were of minor importance for seedling survival on both sites. Safe sites for seed germination and seedling establishment of fen‐meadow species existed on the fen meadow even without soil disturbance and gap creation. Safe sites for seedling recruitment were not present in the rush pasture. The need for species‐specific definition of safe site characteristics at the two stages of seedling recruitment (i.e., for seed germination and for seedling survival) was demonstrated. The implications of these findings for restoration of seminatural grasslands are discussed.     

Community structure and diversity of bryophytes and vascular plants in abandoned fen meadows

We examined effects of abandonment on species diversity and species composition by comparing 21 calcareous fen meadows in the pre-alpine zone of central and northeastern Switzerland. Meadows were divided into three classes of successional stages (mown: annually mown in late summer, young fallow: 4–15 years, and old fallow: >15 years of abandonment). In each fen, we measured litter mass in four 20 cm×20 cm plots, as well as (aboveground) biomass and species density (number of species per unit area) of bryophytes and vascular plants. Bryophyte biomass was reduced in abandoned fens, whereas litter mass and aboveground biomass of vascular plants increased. Species density of both taxonomic groups was lower in abandoned than in mown fens. Young and old successional stages were not different except for bryophytes, for which old successional stages had higher species density than young stages. We used litter mass and aboveground biomass of vascular plants as covariables in analyses of variance to reveal their effects on species density of both taxonomic groups. For bryophytes, litter mass was more important than vascular plant biomass in explaining variance of species density. This indicates severe effects of burying by litter on bryophyte species density. For species density of vascular plants, both vascular plant biomass and litter mass were of similar importance in explaining the decreased species density. Canonical correspondence analyses showed that abandonment also had an effect on species composition of both bryophytes and vascular plants. However, young and old successional stages were not different indicating fast initial changes after abandonment, but slow secondary succession afterwards. Furthermore, indicator species analysis showed that there was no establishment of new species after abandonment that might dramatically alter fen communities. Re-introduction of mowing as a nature conservation strategy may thus be very promising – even for old fallows.     

Comparative ecology of seedling recruitment in an oligotrophic wet meadow

Abstract. For the regeneration niche to contribute to the maintenance of species diversity interspecific differences in sensitivity of seedling recruitment to environmental conditions is assumed. We experimentally tested differences between meadow species for the response of seed germination to chilling, and sensitivity of seedling recruitment to microscale heterogeneity. We also compared the dynamics of seedling recruitment in gaps. Seed germination was tested in standard laboratory germination tests, comparing control seeds with seeds chilled at +4 °C, and at –14°C for one month. Species responses varied from significant increases in germinability after chilling (e.g. Cirsium palustre, Betonica officinalis, Angelica sylvestris) to significant decreases (e.g. Hieracium umbellatum, Succisa pratensis, Selinum carvifolia). In some species, chilling at + 4 °C has a similar effect to chilling at –14 °C, in others the effect of chilling at + 4°C was intermediate, and in some, there was no effect of chilling at + 4°C, but an effect of chilling at –14°C. Different chilling temperatures also affect timing and speed of seed germination under greenhouse conditions. The dynamics of seed germination under field conditions was studied by sowing seeds into artificially created gaps and following their germination, both where the seedlings were removed after emergence and where they were not removed. Species differ in their germination dynamics: they all start late April, but then differentiate from an abrupt maximum and early finish of germination (in the second half of May), to prolonged germination without a marked maximum. Seedling removal increased the total number of germinated seeds, with a marked density dependence at this stage. Seeds were also sown into plots with treatments (1) gaps, sod stripped, above-ground vegetation removed; (2) mown, moss layer removed; (3) mown; (4) untouched control. Seedling emergence was monitored for 3 yr. Seedling recruitment decreased from treatments (1) to (4), but sensitivity differed between species. With increasing seed weight, the difference between gaps and other treatments decreased. The results show that there are considerable differences in seedling recruitment sensitivity between species.     

Dispersal and microsite limitation in an abandoned calcareous grassland of the southern prealps

Dispersal limitation is often involved when the species composition of a dry abandoned grassland shows a slow response to resumed regular mowing. A seed-addition experiment, using 32 species which do not belong to the local species pool, was performed on Monte San Giorgio (southern Switzerland) to test whether the low recruitment success was due to dispersal limitation or due to unfavourable microsite conditions. In October 1997, 20 species were individually sown in six 3 × 4 m blocks of a 2 × 2 factorial “partial” split-plot design with treatments of abandonment vs. mowing and undisturbed vs. root-removed soil, this last being applied in small naturally-degradable pots. Moreover, 12 species were sown only in the treatments on undisturbed soil. Seedlings of sown and spontaneously germinating seeds were observed on 16 occasions over one 12-month period. Seeds of 31 out of the 32 species germinated. Twenty-four species showed germination rates higher than 5% and different seasonal germination patterns. Established vegetation, especially the tussocks ofMolinia arundinacea, reduced the quality of microsites for germination. Whereas a few species germinated better under the litter ofMolinia arundinacea, many more germinated better under the more variable microsite conditions of a mown grassland.     

Plant species decline due to abandonment of meadows cannot easily be reversed by mowing. A case study from the southern Alps

Abstract. We resumed mowing in two plots of ca. 100 m² in an abandoned meadow dominated by Brachypodium pinnatum on the slope of Monte Generoso (Switzerland). We monitored species composition and hay yield using point quadrats and biomass samples. Species frequencies changed little during 10 yr (1988–1997) while hay yields showed large fluctuations according to mean relative humidity in April-June. We performed a seed-addition experiment to test whether the establishment of meadow species is limited by lack of diaspores or favourable microsites for germination and recruitment from the seed bank. We sowed ca. 12 000 seeds of 12 species originating from a nearby meadow individually in plots of a 4 × 6 unbalanced Latin square with four treatments, burning, mowing, mowing and removal of a layer of decayed organic matter, and a control. We monitored the fate of seedling individuals for 24 months. Seedlings of all species were established and survived for 12 months, 10 species survived during at least 24 months, some reached a reproductive stage. Species responded to different qualities of microsites provided by the different treatments thus required different regeneration niches. Spontaneous long-distance immigration was insignificant. We conclude that the former species composition of abandoned meadows cannot easily be restored by mowing alone because many plant species of meadows do not have persistent seed banks and immigration over distances of more than 25 m and successful establishment is very unlikely.     

Short-time effects of land-use changes on O-horizon in subalpine grasslands

Due to economic needs, a process of abandonment of unfavourably situated areas and intensified cultivation on favoured areas has taken place in the alpine region. This abandonment not only means the loss of a cultural landscape based on centuries of care but also interference with an established ecological balance. We wanted to investigate the impact of land-use, such as mowing, fertilisation and irrigation and the effects of abandonment on the O-horizon type, quantity and quality. Therefore, we selected areas with different types of land use but with equivalent site conditions in the Passeier Valley (Bolzano-South Tyrol province) for investigation. The areas studied were as follows: an organically fertilised and irrigated, intensively managed hay meadow mown once a year; an organically fertilised, intensively managed hay meadow mown once a year; a lightly managed hay meadow mown every second year; a lightly managed hay meadow mown every 3–4 years; an area densely covered with dwarf shrubs abandoned 10 years ago; and a reforested area abandoned 30 years ago. These selected areas represent different intensities of land use or stages of succession. On these areas, vegetation, phytomass, litter quantity, humus type, O-horizon quantity, litter decomposition and C and N storage, were analysed in detail. Abandonment causes the percentage of coverage with grasses and herbs to decrease from 100 to 10%, while lignified species increase from 0 to 90%. This leads to growth of the phytomass pools, which in turn causes an increase in the annual litter quantity (from 117 to 525 g m⁻²). Significant changes in litter quality in the case of reduced use or abandonment increases the litter’s resistance to decomposition, decreasing overall decomposition. Litter quantity and quality then have a significant impact on the O-horizon quantity. This increases strongly with decreasing intensity of land use or longer times since abandonment (115–1,180 g m⁻²). This leads to a significant increase of C and N storage and a change in O-horizon type ranging from no definite O-horizon or mull O-horizons on regularly used areas to moder on abandoned areas.     

Microenvironmental conditions for Japanese alder seedling establishment in a hummocky fen

In this article, we focused on hummocky microtopography as a prominent feature of mires and explored the microenvironmental conditions suitable for alder seedling establishment. Japanese alder (Alnus japonica) forest is widely distributed in wetlands in northern Japan. However, because alder seedlings are rare in mires, alder population dynamics and conditions that favor the establishment of alder seedlings are still unknown. The study was conducted in northern Japan at a site in mesotrophic mire. We surveyed the seedling density, the microenvironmental conditions (light, litter cover, and soil quality), and the density of dispersed seeds in alder forest and in the adjacent herbaceous fen. In addition, we performed a laboratory experiment to examine the germination characteristics of alder. Seedlings grew only on hummocks in alder forest. The percentage of litter cover on hummocks was lower than in hollows, and the density of dispersed seeds in alder forest was much higher than in herbaceous fen. Seeds of Japanese alder germinated under both light and dark conditions, and the germination rate were high under light and high-temperature conditions. Our results suggest that litter cover may inhibit seedling establishment and hummocks that characterized by less litter cover are suitable place for the establishment of seedlings. We conclude that hummocky microtopography and abundant seed rain in the mire enable the establishment of Japanese alder seedlings.     

Germination and seedling survival in fens undergoing succession

We studied mechanisms of vegetation change in fens subject to succession from open water to floating mats and finally herbaceous rich-fens. Earlier research showed that these systems are characterized by transient seed banks. Our main question was whether seedlings of later successional fen stages are already present in earlier stages, remaining subordinate in the vegetation until conditions become suitable for them. If, however, conditions during succession change in a way that only a limited set of species can survive as seedlings during each of the successional stages, no seedling bank will exist. The transient character of the seeds would then imply that seeds will not germinate and will subsequently die and that seeds that have germinated in the “wrong” stage will not become established. We hypothesized that: (1) germination and seedling survival of fen species are significantly better in the successional fen stage for which these species are characteristic, (2) as a consequence no seedling banks occur in these fens. In a field experiment, seeds of five characteristic fen species in the standing vegetation of three successional fen stages i.e. raft fen, quaking fen and rich fen were sown in each of these stages in a turf pond in the Tienhoven area, The Netherlands. Germination and seedling survival were measured over two growing seasons together with environmental variables. Germination was higher in the “own” stage for all species groups as was survival for quaking fen species and rich fen species. For both these stages, percentage of germination and survival of four out of five characteristic species were significantly higher in the “own” stage. Germination and survival can be considered stage-dependent and it was concluded that seedling banks do not exist in these fens. Site-specific environmental variables act as a sieve and differentiate on species presence already during early life history stages. We found clues that the environmental sieve acts at the level of nutrient availability, tolerance for high sulphide concentration and light climate. Because of the transient seed bank and absence of a seedling bank in these fen wetlands, successful establishment of species necessitates a continuous dispersal of characteristic species until the environmental conditions permit establishment. This also implies that species of the whole successional sere should be present within dispersal distance.     

Effects of light competition and litter on the performance of Viola palustris and on species composition and diversity of an abandoned fen meadow

Theeffects of light competition and litter on seedling recruitment and theperformance of established individuals were examined in Violapalustris. This polycarpic perennial plant was a common component ofspecies-rich fen meadows in northwestern Germany until the middle of thiscentury, but today is considered to be regionally endangered. From summer 1996until summer 1998 a bi-factorial field experiment combining three standing croptreatments (mowing, thinning, control) with two litter layer treatments (litterremoval, no litter removal) was carried out in an abandoned fen meadow toinvestigate the effects of these factors both on V.palustris and on aboveground species composition and diversity.MANOVAs (multivariate analysis of variance) revealed significant treatmenteffects for the performance of V. palustris related toexperimental manipulation. Mowing increased vital attributes including thenumber of rosette leaves, the percentage of individuals with chasmogamous andcleistogamous flowers and the number of rhizomes. In contrast, the length of theleafstalk, mean leaf area and the length of rhizomes were negatively associatedwith mowing. Litter removal significantly increased the number of V.palustris seedlings. The removal of the litter layer already resultedin an increase in aboveground species richness in the first year of theexperiment. After three years of experimental manipulation, both abovegroundspecies richness and diversity were positively associated with mowing.Redundancy Analysis showed that typical wet meadow species (Cardaminepratensis, Lychnis flos-cuculi, Lotuspedunculatus) and species of mesotrophic fens (Violapalustris, Agrostis canina, Potentillapalustris) were positively influenced by mowing. It can be concludedthat V. palustris recruitment and growth is promoted by aregular disturbance regime on fen meadows. The conservation of many typicalspecies of wet meadows and fens in northwestern Germany depends on moderategrazing or cutting. These management types prevent successional changes whichlead to an increase in standing crop and the development of a litter layer, bothof which can cause local extinction.     

Effects of litter removal and mowing on germination and establishment of two fen-grassland species along a productivity gradient

To study the factors that limit the occurrence of species in fen grasslands, the effects of mowing and litter removal on germination, growth and establishment of two common species (Silene flos-cuculi andLotus pedunculatus) were analyzed along a productivity gradient at seven sites in Northern Germany. The sites differed in nutrient availability, vegetation composition and standing crop. In autumn 2002 a field experiment was set up at each site, including factorial combinations of two treatments (litter removal, mowing). Seeds and juvenile individuals of both species were transferred within the treatment combinations. The number of germinated and transplanted individuals was recorded during the 2003 and 2004 seasons and biomass of survived plants weighed in late summer 2004. Mowing and litter removal had positive effects on the number of germinated seeds of both species. The effect of litter removal was mostly significant at sites with a comparatively low nutrient availability and standing crop (350 g/m²), while the effect of mowing was more important at the site with the highest standing crop (1000 g/m²). In some cases the presence of litter had a positive effect on the survival of individuals, which may be caused by extremely dry weather conditions in 2003. The experiment showed that plant species reaction on mowing and litter removal differed both according to their life stage and the site conditions. These differences could be interpreted in context with Shifting Limitation Hypothesis because the abolition of seed limitation by sowing and creation of additional gaps for germination was most efficient at low productive sites, while the high standing crop reduced first of all the establishment probability, which could be increased by mowing.     

Competitive effects of herbaceous vegetation on tree seedling emergence,growth and survival: Does gap size matter?

Question: What is the effect of gap size on the seedling emergence, growth and survival of four common tree species in wooded pastures? Location: A pasture in the Jura mountains, Switzerland. Methods: Seeds were sown in a complete three‐way factorial design with eight blocks in May 2003. Each block consisted of a competition treatment (four gap sizes including zero) and a mowing treatment (mown and unmown). Emergence, survival and total biomass of tree seedlings of three species (Picea abies, Acer pseudoplatanus and Fagus sylvatica) were measured. A fourth species (Abies alba) failed to germinate. Results: Gaps had a positive influence on the early stages of tree development for all species. Larger gaps favoured growth and survival more than small gaps. Seedling growth was higher when vegetation around the openings was mown. Mowing the vegetation at gap size zero enhanced both growth and survival compared to unmown vegetation. Mown gaps larger than zero had increased seedling desiccation but decreased seedling predation. Species showed similar trends in their emergence and growth responses to gap size and mowing treatments but for Picea emergence rate was higher and survivorship was lower than for Acer and Fagus. Conclusions: Gap size does matter for tree seedling success but even in more favourable large gaps only a small percentage of seedlings emerged and survived. The effects of gaps on tree seedling establishment are complex as a result of interactions between biotic and abiotic changes caused by gaps.     

Effects of litter on seedling establishment: an indoor experiment with short-lived Brassicaceae species

Recruitment by seeds is essential both in vegetation dynamics and in supporting biodiversity in grasslands. The recruitment by seeds is feasible in suitable vegetation gaps from the seed rain and/or by establishment from persistent soil seed banks. Cessation of grassland management results in litter accumulation, which leads to the decline of species diversity by the decreased availability of open patches. Low amounts of litter is often beneficial, while high amounts of litter is detrimental for seed germination and seedling establishment of short-lived species. In a designed indoor experiment, we explored the effect of litter on seedling establishment by germinating six short-lived Brassicaceae species with both increasing seed mass and litter cover. We found that both seed mass and litter had significant effect on germination and establishment of the sown species. Small-seeded species were significantly negatively affected by the 300 and/or 600 g/m² litter layers. No negative litter effect was detected for species with high seed masses (Lepidium spp.). No overall significant positive litter effect was found, although for most of the species cumulative seedling numbers were not the highest at the bare soil pots. Our results suggest that the negative effects of litter are less feasible on the large-seeded short-lived species than on that of small-seeded ones.     

Regeneration by seeds in alpine meadow and heath vegetation in sub‐arctic Finland

Abstract. This paper compares the regeneration by seeds of heath and meadow and studies relationships between the floristic composition of phases in the regeneration pathway. Seed densities in the seed rain and seed bank as well as the densities of emerged seedlings in gaps and in closed vegetation were greater in the meadow than in the heath. In the heath, environmental constraints hindered seedling emergence almost completely so seeds accumulated in the seed bank. In the meadow, the decrease in the seed bank was due to high seedling emergence. Within both plant communities, seedling emergence in gaps and in closed vegetation was comparable. In the meadow, the seed rain and seedling emergence in gaps, as well as the seed bank and seedling emergence in gaps were positively correlated. Differences in seed and adult plant sizes were reasons for the low correlation between the standing vegetation and the other phases. In DCA ordination the first axis separated the phase of seedling emergence in closed vegetation and seed bank. The second axis separated the standing vegetation from the other phases. The structure of the seed rain was more heterogeneous than that of other phases. In the heath, the standing vegetation and the seed rain were positively correlated. The ordination of these phases reflected the patchiness of standing vegetation and the ability of the diaspores of Betula nana to disperse over long distances.     

Mechanisms of structural change derived from patterns of seedling emergence and mortality in a semi-natural meadow

Questions: Is seedling emergence limited by the set of viable seeds, by incompatibility between the phenology of seed shedding and timing of mowing, or by dry weather in germination periods? Does seedling mortality fluctuate with season and weather? Location: Negrentino, southern Alps, Switzerland. Methods: Fecundity estimates of the dominant grass Bromus erectus; highly frequent counts of spontaneous seedlings by species and calculation of a community-level average mortality rate across 5 years; species-level records of seed shedding date and measurements of seed mass; measurement of soil moisture. Results: B. erectus produced 143.9 viable seeds/m²/year while the density of its seedlings was a 55 times smaller fraction. Grasses had fewer seedlings than forbs and their phenology of seed shedding was less compatible with mowing date. Soil moisture was a strong determinant of seedling emergence in spring and less so in autumn. Average seedling mortality declined with age of the populations and reached a maximum in an extremely dry summer. In relatively wet summers establishment success was positively related to seed mass. Conclusion: Community structure is susceptible to drought through mechanisms that selectively reduce recruits of coexisting plant functional groups. We propose that (1) more frequent intense droughts tend to reduce species that depend on frequent recruitment from seed, hence favour long-lived clonally spreading species, (2) drought timing selects between species with different germination phenology and drought resistance, and (3) drought impacts can be mitigated by changing management regimes that affect seed shedding.